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Habitat Restoration Opportunities, Climatic Niche Contraction, and Conservation Biogeography in California's San Joaquin Desert

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Habitat Restoration Opportunities, Climatic Niche Contraction, and Conservation Biogeography in California's San Joaquin Desert Habitat restoration opportunities, climatic niche contraction, and conservation biogeography in California's San Joaquin Desert Running head: Habitat restoration opportunities and climatic niche contraction in California's San Joaquin Desert Joseph A E Stewart1,2*, H Scott Butterfield3, Jonathan Q Richmond4, David J Germano5, Michael F Westphal6, Erin N Tennant7, Barry Sinervo1,2 1 Department of Ecology and Evolutionary Biology, University of California Santa Cruz, 1156 High St, Santa Cruz, CA 95064, USA. 2 Institute for the Study of Ecological and Evolutionary Climate Impacts, University of California, 100 Shaffer Road, Santa Cruz, CA 95060, USA. 3 The Nature Conservancy, 201 Mission St, San Francisco, CA, 94105, USA. 4 U.S. Geological Survey, 4165 Spruance Rd., Suite 200, San Diego, CA, 92101, USA. 5 Department of Biology, California State University Bakersfield, 9001 Stockdale Hwy, Bakersfield, CA, 93311, USA. 6 U.S. Bureau of Land Management, 940 2nd Ave., Marina, CA, 93933, USA. 7 Lands Unit, Central Region, California Department of Fish and Wildlife, 1234 E. Shaw Ave., Fresno, CA, 93710, USA. * To whom correspondence may be addressed. E-mail: [email protected] 1 PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.26758v2 | CC0 Open Access | rec: 10 Jan 2019, publ: 10 Jan 2019 Abstract A recent global trend toward retirement of farmland presents opportunities to reclaim habitat for threatened and endangered species. We examine habitat restoration opportunities in one of the world’s most converted landscapes, California’s San Joaquin Desert (SJD). Despite the presence of 35 threatened and endangered species, agricultural expansion continues to drive habitat loss in the SJD, even as marginal farmland is retired. Over the next decades a combination of factors, including salinization, climate change, and historical groundwater overdraft, are projected to lead to the retirement of more than 2,000 km2 of farmland in the SJD. To promote strategic habitat protection and restoration, we conducted a quantitative assessment of habitat loss and fragmentation, habitat suitability, climatic niche stability, climate change impacts, habitat protection, and reintroduction opportunities for an umbrella species of the SJD, the endangered blunt-nosed leopard lizard (Gambelia sila). We use our suitability models, in conjunction with modern and historical land use maps, to estimate the historical and modern rate of habitat loss to development. The estimated amount of habitat lost since the species became protected under endangered species law in 1967 is greater than the total amount of habitat currently protected through public ownership and conservation easement. We document climatic niche contraction and associated range contraction away from the more mesic margins of the species’ historical distribution, driven by the anthropogenic introduction of exotic grasses and forbs. The impact of exotic species on G. sila range dynamics appears to be still unfolding. Finally, we use NASA fallowed area maps to identify 610 km2 of fallowed or retired agricultural land with high potential to again serve as habitat. We discuss conservation strategies in light of the potential for habitat restoration and multiple drivers of ongoing and historical habitat loss. Keywords: climatic niche shift, extirpation, farmland retirement, invasive species, range contraction, reintroduction Introduction Habitat loss resulting from agricultural expansion is one of the greatest historical drivers of extinction [1,2]. A recent global trend toward retirement of marginal farmland, especially in temperate latitudes, presents an important opportunity to reclaim some of this lost habitat in a cost-effective manner before the land is claimed for other uses [3–6]. More information on patterns of habitat loss, degradation, and use will allow targeted habitat restoration in areas where it will make the biggest impact for conserving sensitive and endangered species. To better understand this opportunity, we investigate habitat and land use trends in the San Joaquin Desert (SJD) of California [7]. Now largely converted to agriculture, the SJD was once an extensive network of upland and aquatic habitats, supporting a high concentration of endemic species [8]. As a result of habitat loss, 7 species have been driven to extinction or regional extirpation from the SJD, and 35 species are now protected under endangered species legislation (S1 Table). Over the next 30 years a combination of factors, including salinization, climate change, and historical groundwater overdraft, are projected to lead to the retirement of more than 2,000 km2 of SJD farmland [9]. If retired farmland can be restored to habitat in strategically selected areas of the SJD, it could contribute to the recovery of dozens of vulnerable species. While we center our discussion primarily around restoration opportunity, which has great potential to benefit many endangered species, we also present documentation of a rarely 2 PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.26758v2 | CC0 Open Access | rec: 10 Jan 2019, publ: 10 Jan 2019 documented phenomenon: climatic niche contraction mediated by invasive species. Previous studies documented negative impacts of exotic grasses and forbs on species demography [10,11]. Here we document climatic niche contraction and associated range contraction away from the mesic margin of a species historical distribution. The contraction was driven by the interaction of precipitation and invasive grasses and forbs. Though we suspect the phenomenon is widespread, this study appears to be one of few empirical examples of climatic niche contraction caused by an invasive species (but see [12]). The phenomenon has interesting implications for community reshuffling in the context of climate change. In this study we focus on habitat change in an iconic endangered species of the SJD, the blunt-nosed leopard lizard (Gambelia sila), known for its large size, bright nuptial coloration and bipedal sprint bursts [13]. We focus on G. sila in part because the habitat requirements of the species are generally representative of other upland species of the SJD [7,14]. We use modern and historical land use maps to estimate historical and modern rates of habitat loss and to assess habitat fragmentation. We use historical occurrence records, resurveys, and vegetation measurements to document a changing climatic niche. We also use NASA fallowed area maps to identify 610 km2 of fallowed or retired agricultural land with high potential to be restored as habitat. We build habitat suitability models that incorporate known ecophysiological mechanisms that govern species distributions [15–17]. Unlike previous models of habitat suitability for species of the SJD [6,18] (but see [19], S2 Table), our models are appropriate for assessing habitat restoration potential on retired farmland because they properly account for anthropogenic land use. We discuss conservation strategies in light of the potential for habitat restoration and multiple drivers of ongoing and historical habitat loss. Methods Study system The San Joaquin Desert (SJD) of California encompasses 28,493 km2 including the western and southern two-thirds of the San Joaquin Valley, as well as the Carrizo Plain and Cuyama Valley to the southwest [7]. The SJD is distinguished from the larger San Joaquin Valley by low average precipitation ( 279 mm annually), aridic soils, and the presence of a high concentration of co-occurring endemic plant and animal species [8]. The SJD once supported extensive upland habitat composed of alkali sink scrub, saltbush shrub (Atriplex spp.), Ephedra scrubland, and grassland dominated communities as well as a vast aquatic system of lakes, rivers, marshes, and sloughs fed by rainfall and snowmelt from the Sierra Nevada. Today, most native habitat has been converted to row crops and orchards, remnant upland habitat is heavily impacted by exotic annual grasses and forbs, and wetlands have been drained to support agriculture [20–22]. Thirty-five threatened or endangered species are now confined to isolated patches of habitat in the SJD (S1 Table). We focus our analyses on habitat of an endemic species of the SJD, G. sila. Gambelia sila were among the first species protected under United States endangered species legislation in 1967 [23] and remain listed as endangered today. Their status is mainly a result of habitat loss and fragmentation, energy development, and non-native vegetation [24]. They use their powerful hind limbs to sprint while evading predators and while catching prey, which consist largely of 3 PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.26758v2 | CC0 Open Access | rec: 10 Jan 2019, publ: 10 Jan 2019 coleopterans and orthopterans [25]. Gambelia sila inhabit relatively flat, sparsely vegetated areas of the SJD including the valley floor, surrounding foothills, and valleys to the southwest [7,8,26]. Occurrence data We used 618 geographically unique records of G. sila occurrence to develop habitat suitability models. We obtained occurrence data from publicly available data portals (e.g., VertNet.org, GBIF.org), the California Natural Diversity Database (CNDDB), correspondence with professional biologists, the literature, and from surveys conducted by the authors of this paper. We corrected for sample bias [27–29] by using all geographically unique vertebrate occurrence records within 50 km of occurrence locations as background or pseudo-absence data (n = 6,285). We did not consider background
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